Medical and Veterinary Entomology (2011), doi: 10.1111/j.1365-2915.2010.00939.

Variations in life tables of geographically isolated

strains of the mosquito Culex quinquefasciatus
1

D. S. S U M A N , S. N. T I K A R , M. J. M E N D K I , D. S U K U M A R A N ,
2
1
1
O. P. A G R A W A L , B. D. P A R A S H A R and S. P R A K A S H
1 Division

of Entomology, Defence Research and Development Establishment, Gwalior, India and

Jiwaji University, Gwalior, India

2 School

of Studies in Zoology,

Abstract. Variations in the life tables and other biological attributes of four strains of
Culex quinquefasciatus Say (Diptera: Culicidae) from geographically isolated regions
of India that had been reared to the fifth generation in the laboratory were assessed
under a standardized rearing regime under constant laboratory conditions. Two strains
from arid habitats [Jodhpur (JD) and Bikaner (BKN)], one from a semi-arid inland
habitat [Bathinda (BTH)], one from a semi-arid coastal habitat [Jamnagar (JMN)] and
a standard laboratory strain (LAB) were compared. Horizontal life-table parameters
were measured for each strain. Egg mortality ranged from 4.4% (JD and BTH) to
19.5% (BKN). The lowest rate of adult emergence and highest female : male ratio
were found in BKN, and the highest rate of adult emergence and lowest female : male
ratio were recorded in BTH. The egg-hatching period was longest in BTH and shortest
in LAB. The duration from oviposition to adult emergence was longest in JD and
shortest in LAB. Females lived longer than males in all strains. The net reproductive
rates (R0 ) of all field-derived strains (122.9162.2) differed significantly between
strains and were significantly greater than that of LAB (107.6). Similarly, both the
intrinsic rate of increase (rm ) and finite rate of increase () were found to be lower
in LAB than in the field strains, but the mean generation time (T ) and doubling time
(DT ) were longest in LAB. For several life-table attributes, JD and BTH clustered
together and were more similar to JMN than to BKN and LAB. The results indicate
that BTH, BKN and JD can be characterized as r-strategists, more so than JMN.
Overall fecundity increased with age. Differences in annual temperature ranges and
mean annual rainfall between locations were positively correlated (r = 0.460.97)
with egg production, female life expectancy, R0 , rm , and T . The results suggest that
strains of Cx. quinquefasciatus from different geographical areas with contrasting
habitats vary in their survival and reproductive strategies accordingly.
Key words. Culex quinquefasciatus, ecological variation, life reproductive strategies,

life-table attributes, strain development.

Introduction
Culex quinquefasciatus Say is the most common domestic
species of mosquito in the tropics. It is the principal vector of
Bancroftian filariasis (World Health Organization, 2006) and

breeds in diverse aquatic habitats, ranging from polluted to

fresh water (Sirivanakaran, 1976).
The survival and fecundity of different mosquito species are
affected by factors such as nutrition (Day et al., 1994; Costero
et al., 1998), resistance to organophosphate insecticides (Jinfu

Correspondence: Dr B. D. Parashar, Scientist F, Division of Entomology, Defence Research and Development Establishment, Gwalior, Madhya
Pradesh 474002, India. Tel.: +91 751 223 1862; Fax: +91 751 234 1148; E-mail: bdparashar@rediffmail.com
2011 The Authors
Medical and Veterinary Entomology 2011 The Royal Entomological Society

2 D. S. Suman et al.
et al., 1998), temperature (Maharaj, 2003) and infection by
microbial pathogens (Mahmood et al., 2004). Mosquitoes
occurring in distinct locations normally adapt to local conditions. Over time, ecological isolation and adaptation to local
conditions may lead to the development of geographically isolated strains that differ in various biological traits, such as
fecundity and survivorship, which gives rise to variation in a
range of population parameters (Reisen et al., 1979).
Distinctive life-table attributes have been found in strains
of Cx. quinquefasciatus from Bangkok (Thailand), the southeastern U.S.A. (Vero Beach, FL), the southwestern U.S.A.
(Houston, TX) (Walter & Hacker, 1974) and the Northwest
Frontier in Pakistan (Suleman & Reisen, 1979). Recently,
Suman et al. (2009) reported significant variations in morphometrics and morphology among eggs from the four strains of
Cx. quinquefasciatus investigated in the present study, sourced
from arid, semi-arid and coastal regions of India. We hypothesized that these mosquito strains will differ in their life-table
and biological traits because they appear to be adapted to their

respective local environments. To test this notion, we assessed

life-table and other biological attributes under standardized laboratory environments in four strains of Cx. quinquefasciatus
from a range of distinct isolated habitats in India and a standard
laboratory strain, and correlated the attributes with ecological
variables.

Materials and methods

Mosquito collection, rearing and experimentation
Culex quinquefasciatus larvae were collected from military
cantonments in Jodhpur and Bikaner on the western plain of an
arid zone in the Thar Desert, an air force station in Jamnagar
on the west coastal plain of a semi-arid zone close to the Arabian Sea coast, and Bathinda on a northern plain of the upper
northwestern region of a semi-arid zone (Fig. 1). The annual
range of temperature (winter minimumsummer maximum)

km

Fig. 1. Locations of collections of Culex quinquefasciatus in India. BTH, Bathinda; BKN, Bikaner; JD, Jodhpur; JMN, Jamnagar; GWL, laboratory
at Gwalior.
2011 The Authors
Medical and Veterinary Entomology 2011 The Royal Entomological Society, Medical and Veterinary Entomology, doi: 10.1111/j.1365-2915.2010.00939.x

Culex quinquefasciatus life tables 3

was intermediate in Jodhpur and Bikaner (245 C), mildest
in Jamnagar (936 C) and most extreme in Bathinda
(247 C). Annual rainfall ranged from <200 mm in
Bikaner, to 200500 mm in Jodhpur and Bathinda and
5001000 mm in Jamnagar. Although Jodhpur receives more
rainfall than Bikaner, both have desert vegetation. Coastal Jamnagar differs from the other sites in that it has the least extreme
range of temperatures and the highest rainfall, and features
tropical thorn and littoral or swampy vegetation. Bathinda has
the greatest range of temperatures and relatively moderate rainfall, and has tropical dry deciduous vegetation. The laboratory
strain originated in Gwalior, Madhya Pradesh and has been
colonized in our laboratory for >10 years. This is situated in
a semi-arid region that receives 5001000 mm of rainfall per
year. Annual temperature ranges between 2 C and 48 C. The
vegetation of this region is tropical deciduous and thorny.
Larvae were brought to the Defence Research and Development Establishment, Gwalior and maintained in standard
laboratory conditions at 27 1 C, 75 5% relative humidity (RH) and light : dark (LD) 12 : 12 h. Larvae were kept at
a density of 100 larvae per bowl, in enamel bowls containing 2 L of dechlorinated water and were fed regular amounts
of dried yeast on alternate days. After pupation, they were
transferred to cages (75 65 65 cm) to provide up to 50
male and 50 female adults per cage after emergence. After
emergence, adults were provided with 10% sugar solution ad
libitum dispensed through a cotton wick in a Petri dish and
a chick was provided as a source of blood twice a week.
These colonies were maintained for up to four generations
(approximately 2 months).Chicks were maintained in accordance with approved guidelines of the Committee for the Purpose of Control and Supervision of Experiments on Animals
(CPCSEA), India.
Although laboratory rearing imposes a certain degree of
selection pressure on aspects of mosquito biology, all colonies
were exposed to standardized environmental conditions that
were favourable to mosquito survival; hence, we assume that
estimates of life-table parameters derived from data collected
from the colonized wild strains represent a maximum expression of their life-table parameters and are likely to reflect true
differences between geographically isolated strains. Similar
assumptions were made by Reisen et al. (1979) to compare
the life-table strategies of geographically distinct strains of
Culex tritaeniorhynchus after colonization in standard laboratory conditions for many generations (range: 63236).
To assess the adult parameters, 50 pairs of male and female
mosquitoes of the same age from the F5 generation were
released in each of three rearing cages for each strain [i.e.
four experimental strains from Jodhpur (JD), Bikaner (BKN),
Bathinda (BTH) and Jamnagar (JMN), respectively, plus the
laboratory strain (LAB)]. Adults were provided with 10% sugar
solution and a chick for blood feeding ad libitum throughout
the experiment. A glass Petri dish (140 mm in diameter)
containing 400 mL water was provided for oviposition. Daily
observations of egg rafts laid and the mortality or survival of
adult male and female mosquitoes were recorded until the death
of all mosquitoes. The numbers of egg rafts laid were counted,
after which the numbers of eggs per raft were counted under

a microscope (Lieca MZ12; Leica Microsystems, Wetzlar,

Germany). For each strain, the three largest egg rafts were
selected and placed individually into three bowls, each
containing 2 L of dechlorinated water. Subsequently, various
developmental parameters of immature stages were recorded
for each strain. Hatching rates were calculated by comparing
the number of eggs per egg raft with the number of larvae
produced 12 h later. This was carried out by observing the
larvae against a contrasting background and removing them
by pipette to a new bowl. Egg rafts with unhatched eggs
were transferred to separate bowls for further hatching and
counted at 24-h intervals until hatching of larvae was complete;
numbers of late-hatching larvae were added to the initial count.

Statistical analysis of life-table and other biological attributes

Data analysis was based on mean values for the three replicated cohorts of 50 females for each strain. Data were analysed for mean pre-oviposition days (mean number of days
from first exposure to blood host to the appearance of the
first egg raft), oviposition days (mean number of days of
egg laying) and post-oviposition days (mean number of days
from the last egg raft laid to the death of the last female),
mean number of eggs produced per cohort of 50 females,
mean number of eggs per female lifespan [mean number
of eggs/female/oviposition (i.e. number of eggs/number of
females alive on each oviposition day) for all oviposition days
until all females were dead number of oviposition days]
and eggs/female/day (eggs/female lifespan/number of oviposition days). Various attributes of a horizontal life table were
estimated, including life expectancy at emergence (ex ), net
reproductive rate (R0 ), intrinsic rate of increase (rm ), finite
rate of increase () and mean generation time (T ) as per the
method of Howe (1953), Andrewartha & Birch (1954) and
Slobodkin (1962). Population doubling time (DT ) was also
calculated (Yurttas & Alten, 2006). The equations used for
these calculations were as follows:
1 age-specific survivorship: lx = yx /y0 , where yx = the
number of males or females alive on each day, x;
2 life
w expectancy at emergence: ex = Tx / lx , where Tx =
x=1 Lx and Lx = (lx + lx +1 )/2, where w = the day on
which the last individual died;
3 net reproductive rate: R0 = w
x=1 lx mx , where mx =
mean number of eggs/2 (i.e. assuming a 1 : 1 sex ratio
for progeny, the mean number of eggs is divided by 2 to
give the number of female eggs;

4 mean generation time: T = W
X=1 xlx mx /R0 ;
5 intrinsic rate of increase: rm = log e R0 /T , where log e
is the natural logarithm;
6 finite rate of increase: = anti-log erm , and
7 doubling time: DT = (loge2 )/rm.
Percentages of apparent mortality of each life stage (percentage of dead individuals out of the total number of individuals
that entered that stage) and actual mortality (percentage of
dead individuals in a particular stage out of the total number of individuals at the beginning of that generation) for each

2011 The Authors

Medical and Veterinary Entomology 2011 The Royal Entomological Society, Medical and Veterinary Entomology, doi: 10.1111/j.1365-2915.2010.00939.x

4 D. S. Suman et al.
developmental stage (i.e. eggs, larvae, pupae and adults) were
calculated (Southwood, 1978). The sex ratio (female : male)
(Maharaj, 2003), egg-hatching period and periods from egg
laying to development of larvae, pupae and adults were also
calculated.
In order to compare findings among all the strains, all the
biological and life-table parameters of these strains were subjected to analysis of variance (anova) to obtain the least
significant difference (LSD) at P < 0.05. Regression analysis was conducted to reveal the relationship between fecundity
(Y-function; i.e. eggs/female, egg rafts/female and eggs/egg
raft) and age of female mosquitoes (X-function). The range of
annual temperature (difference between minimum winter and
maximum summer temperatures) and the mean annual rainfall for the study areas were correlated with various life-table
attributes. Cluster analysis with the nearest neighbour method
and the squared Euclidean distance metric method (Krebs,
1989) (statgraphics Plus 5.0; StatPoint Technologies, Inc.,
Warrenton, VA, U.S.A.) was used to assess degrees of similarity among different strains according to 14 important life-table
and biological attributes of immature and adult stages, including actual mortality (percentages) of eggs, larvae and pupae,
rate of adult emergence, female : male ratio, total egg production by cohort, eggs laid per female per day, life expectancies
of males and females, R0 , rm , , T and DT. For the analysis
of r-strategy, the ex of females, and the rm , and DT of all
strains including LAB, were considered.

The rank order of the various strains by percentage of actual

mortality in the larval stages was JMN < JD < LAB < BTH
< BKN (Table 1). Statistically significant differences in percentages of both apparent and actual mortality were observed
among LAB, BKN and JMN. However, LAB did not show
significant differences from JD and BTH for either parameter.
Differences between JMN and BTH were significant for the
percentage of actual mortality, but not for the percentage of
apparent mortality.
The highest percentages for actual and apparent pupal mortality were recorded in JD and the lowest in BTH (Table 1).
Both of these strains differed significantly from the other
strains in this respect.
The rank order of the various strains for adult emergence
was BKN < LAB < JD < JMN < BTH; rates ranged from
42.0% to 74.0% (Table 1). The female : male ratio was >1.0
in all strains except BTH (Table 1).

Developmental period for different stages

of Cx. quinquefasciatus
Only LAB and BTH differed significantly from one another
on length of egg-hatching period. The minimum and maximum
periods from egg laying to the development of pupae occurred
in BKN and JD, respectively. The minimum and maximum
periods from egg laying to adult emergence occurred in LAB
and JD, respectively (Table 1).

Results
Mortality in different stages of immature Cx. quinquefasciatus

Survival of adult mosquitoes

The actual and apparent percentages of egg mortality

(Table 1) were highest in BKN (19.5%) and lowest in JD
and BTH (4.4%). On these parameters, JD and BTH differed
significantly from LAB, BKN and JMN. The LAB strain
differed significantly from BKN, but not from JMN.

Overall, females lived longer than males (Figs 2 and 3,

Table 2). Life expectancies at emergence (ex ) of males differed significantly among all strains, except between JMN and
JD, and BTH and LAB. In females, ex differed significantly
among all strains, except between BKN and LAB.

Table 1. Mortality rates and other developmental parameters in immature Culex quinquefasciatus of four strains from different geographical areas
and a strain raised under laboratory conditions.

Parameters
Eggs hatching, %
Mortality rates
Eggs
Larvae
Pupae
Developmental parameters
Egg-hatching period, days
Eggpupa period, days
Eggadult period, days
% adult emergence
Sex ratio (female : male)

%
%
%
%
%

actual and apparent

actual
apparent
actual
apparent

LAB strain,
mean SE

JD strain,
mean SE

BKN strain,
mean SE

JMN strain,
mean SE

BTH strain,
Mean SE

LSD

86.7

95.6

80.5

85.9

95.6

2.10b

7.50

1.21a

0.55b

2.40a

8.50a

13.3 1.21a
14.5 0.60a
16.7 0.71a
7.7 0.43a
10.6 0.49a

4.4 0.55b
13.0 1.13a
13.6 1.25ac
14.8 0.59b
17.9 1.68b

19.5 2.37c
32.3 2.89b
40.1 4.23b
6.3 0.88a
13.1 2.16a

18.1 3.90ac
6.6 1.02c
8.1 1.65c
7.0 1.18a
9.4 2.04a

4.4 2.10b
17.2 3.05a
15.2 6.94ac
4.4 0.48ac
5.0 1.62c

5.28
4.39
8.08
1.83
3.93

1.5 0.22a
10.0 0.71ab
11.6 0.22a
64.5 1.05a
1.1 0.05a

1.8 0.51ab
11.8 0.51b
14.8 0.51b
67.7 1.57ac
1.0 0.04ac

1.7 0.1ab
8.8 1.50a
11.7 0.58a
42.0 4.46b
1.2 0.06b

2.0 0.36ab
9.5 1.52ab
12.0 0.36a
68.3 5.88a
1.1 0.06a

2.4 0.58b
9.3 0.51ab
12.2 0.36a
74.0 5.56c
1.0 0.03c

0.87
2.55
0.97
9.33
0.12

Mean values within rows indicated by same letter do not differ significantly.
LAB, laboratory; JD, Jodhpur; BKN, Bikaner; JMN, Jamnagar; BTH, Bathinda; SE, standard error; LSD, least significant difference.
2011 The Authors
Medical and Veterinary Entomology 2011 The Royal Entomological Society, Medical and Veterinary Entomology, doi: 10.1111/j.1365-2915.2010.00939.x

Culex quinquefasciatus life tables 5

Fecundity

1
0.9
0.8
0.7

lx

0.6
0.5
0.4
0.3
0.2
0.1
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61
Days
Lab

JD

BKN

JMN

BTH

Fig. 2. Survival of males in the LAB, JD, BKN, JMN and BTH strains
of Culex quinquefasciatus.
1
0.9
0.8
0.7

The rank order of the various strains for egg production

by cohort of 50 females was LAB < BKN < BTH < JD
< JMN, with significant differences among strains (Table 2).
The highest and lowest numbers of egg rafts were produced by
JMN and LAB, respectively. Significant differences in egg raft
production were observed among all strains, except between
BTH and LAB. The number of eggs per egg raft was highest
in BTH (n = 207.5) and lowest in JMN (n = 161.1), with
significant differences among all field strains, but not between
LAB and BKN or JMN. The largest (mean 317.3 eggs/raft) and
the smallest (mean 73.0 eggs/raft) were recorded in BKN. No
significant differences were recorded among all field strains for
largest and smallest egg rafts, except for BTH for the smallest
egg rafts. The number of eggs/female lifespan was greatest in
BTH (812.3 eggs) and least in LAB (512.4 eggs), and varied
significantly among strains. Eggs/female/day ranged from 20.5
(LAB) to 30.8 (BTH), showing some significant differences
between strains.

lx

0.6
0.5

Effect of ageing on fecundity

0.4
0.3

Regression slopes indicate that the mean number of egg rafts

per female (Table 3, Fig. 4) and the mean number of eggs per
female (Table 4, Fig. 5) increased significantly with increasing
age of females in all strains except JD. Reductions in the mean
number of eggs per raft with increasing age of females were
variable across the strains (Table 5, Fig. 6).

0.2
0.1
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61
Days
Lab

JD

BKN

JMN

BTH

Fig. 3. Survival of females in the LAB, JD, BKN, JMN and BTH
strains of Culex quinquefasciatus.

Oviposition schedule
Only JD differed significantly from the rest of the strains in
oviposition days. No significant differences were recorded for
pre- and post-oviposition days in any strain (Table 2).

Adult life-table attributes

All strains differed significantly from one another in the
net reproductive rate (R0 ) (Fig. 7), with the maximum occurring in JMN (162.2) and the minimum in LAB (107.6). The
intrinsic (rm ) and finite () rates of increase were least in
LAB and greatest in BTH, with significant differences between

Table 2. Comparison of biological attributes in Culex quinquefasciatus of four strains from different geographical areas and a strain raised under
laboratory conditions.

Parameters

LAB strain,
mean SE

JD strain,
mean SE

BKN strain,
mean SE

JMN strain,
mean SE

BTH strain,
mean SE

LSD

Pre-oviposition, days
7.0
6.3
6.7
8.0
7.3
2.47
Oviposition, days
25.3 0.58a
21.0 1.00b
25.7 1.15a
24.7 2.08a
26.3 0.58a
2.67
Post-oviposition, days
7.3 0.58a
6.7 1.15a
4.7 0.58a
4.0 2.65a
5.3 0.58a
3.42
Total egg raft production by cohort
63.4 2.08a
80.0 1.00b
74.3 3.51c
100.7 2.89d
65.3 0.58a
5.45
Largest egg raft
276.3 35.80a 308.0 20.07ab 317.3 19.14b 287.7 2.31ab 310.3 8.08b 32.52
Smallest egg raft
83.7 3.51a
74.7 0.58ab
73.0 5.20b
77.0 1.73ab
95.3 5.77c
9.82
Eggs per egg raft
169.1 2.99ab
186.3 3.49c
170.2 5.16a
161.1 2.64b
207.5 0.92d
8.03
Total egg production/cohort
10 763 196a
14 903 320b
12 639 213c
16 218 196d 13 560 129e 533
Eggs/female/day
20.5 0.26a
28.2 0.77b
20.5 1.91a
27.9 2.13b
30.8 0.50c
2.38
Eggs/female lifespan
512.4 6.64a
573.9 1.67b
552.4 12.80c 684.3 3.67d
812.3 4.66e 17.39
Life expectancy (ex ) (at emergence), days
21.2 0.15a
19.2 0.21b
18.2 0.38c
19.8 1.02b
21.3 0.40a
0.97
Life expectancy (ex ) (at emergence), days
27.4 0.16a
30.8 0.28b
28.0 0.40a
36.9 0.70c
25.2 0.60d
0.81
1.00a

0.58a

0.58a

1.73a

0.58a

Mean values within rows indicated by same letter do not differ significantly.
LAB, laboratory; JD, Jodhpur; BKN, Bikaner; JMN, Jamnagar; BTH, Bathinda; SE, standard error; LSD, least significant difference.
2011 The Authors
Medical and Veterinary Entomology 2011 The Royal Entomological Society, Medical and Veterinary Entomology, doi: 10.1111/j.1365-2915.2010.00939.x

6 D. S. Suman et al.
Table 3. Regression analysis for number of egg rafts/female (Y) as a function of age (X) of females in Culex quinquefasciatus of four strains
from different geographical areas and a strain raised under laboratory conditions.
Strain

Source of variation

Mean square

Regression equation

d.f.

F-ratio

P -value

LAB

Regression
Residual
Regression
Residual
Regression
Residual
Regression
Residual
Regression
Residual

0.0424
0.00335
0.00848
0.00354
0.07664
0.00609
0.13987
0.00626
0.20294
0.00931

Y = 0.0140 + 0.0026X

1
34
1
29
1
34
1
31
1
31

12.67

0.0011

2.39

0.1327

12.58

0.0012

22.33

0.0001

21.79

0.0001

JD
BKN
JMN
BTH

Y = 0.0665 + 0.0011X
Y = 0.0002 + 0.0038X
Y = 0.00588 + 0.004X
Y = 0.06088 + 0.0072X

LAB, laboratory; JD, Jodhpur; BKN, Bikaner; JMN, Jamnagar; BTH, Bathinda.
Jodhpur strain JD

Laboratory strain LAB

0.24
Y = 0.0140 + 0.0026X

Egg raft/female

Egg rafts/female

0.25
0.2
0.15
0.1
0.05

Y= 0.0666 + 0.0011X

0.2
0.16
0.12
0.08
0.04

0
0

10

20

30

40

50

60

Age of female, Days

(A)

30

40

50

60

Jamnagar strain JMN

0.5

0.4

Y = -0.0059 + 0.004X

Y = 0.002 + 0.0038X

Egg rafts/female

Egg raft/female

20

Age of female, Days

(B)

Bikaner strain BKN

0.3
0.2
0.1
0
0

(C)

10

10

20

30

40

0.4
0.3
0.2
0.1
0

50

Age of female, Days

20

40

60

80

Age of female, Days

(D)

Bathinda strain BTH

Egg rafts/female

0.6
Y = -0.061 + 0.0072X

0.5
0.4
0.3
0.2
0.1
0
0

(E)

10

20

30

40

50

Age of female, Days

Fig. 4. Regression line between egg rafts per female and age of female (days) in Culex quinquefasciatus in (A) LAB, (B) JD, (C) BKN, (D) JMN
and (E) BTH strains.
2011 The Authors
Medical and Veterinary Entomology 2011 The Royal Entomological Society, Medical and Veterinary Entomology, doi: 10.1111/j.1365-2915.2010.00939.x

Culex quinquefasciatus life tables 7

Table 4. Regression analysis for number of eggs/female (Y) as a function of age (X) of females in Culex quinquefasciatus of four strains from
different geographical areas and a strain raised under laboratory conditions.
Strain

Source of variation

LAB

Regression
Residual
Regression
Residual
Regression
Residual
Regression
Residual
Regression
Residual

JD
BKN
JMN
BTH

Mean square
594.577
115.374
142.008
133.205
840.04
138.514
2328.9
160.481
7102.28
453.736

Regression equation

d.f.

F-ratio

Y = 5.7396 + 0.3168X

1
33
1
28
1
34
1
31
1
31

5.15

0.029

1.07

0.3107

6.06

0.019

14.51

0.0006

15.65

0.0004

Y = 13.8519 + 0.1612X
Y = 4.6648 + 0.4075X
Y = 2.5433 + 0.5467X
Y = 9.720 + 1.3603X

P -value

LAB, laboratory; JD, Jodhpur; BKN, Bikaner; JMN, Jamnagar; BTH, Bathinda.
Jodhpur strain JD

Laboratory strain LAB

50

50

Eggs/female

Eggs/female

Y= 5.7396 + 0.3168X

40
30
20

Y = 13.851 + 0.1612X

40
30
20
10

10

0
0

10

(A)

20

30

40

50

60

10

Age of female, Days

30

40

50

60

Age of female, Days

Jamnagar strain JMN

Bikaner strain BKN

80

60

Y = 2.5433 + 0.5467X

Y = 4.6648 + 0.4075X

Eggs/female

50

Eggs/female

20

(B)

40
30
20

60
40
20

10

0
0
(C)

10

20

30

40

50

20

(D)

Age of female, Days

40

60

80

Age of female, Days

Bathinda strain BTH

150

Egg/ female

Y = -9.720 + 0.1.360X

120
90
60
30
0
0

(E)

10

20

30

40

50

Age of female, Days

Fig. 5. Regression line between eggs per female and age of female (days) in Culex quinquefasciatus in (A) LAB, (B) JD, (C) BKN, (D) JMN and
(E) BTH strains.
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8 D. S. Suman et al.
Table 5. Regression analysis for number of eggs/egg raft (Y) as a function of age (X) of females in Culex quinquefasciatus of four strains from
different geographical areas and a strain raised under laboratory conditions.
Strain

Source of variation

LAB

Regression
Residual
Regression
Residual
Regression
Residual
Regression
Residual
Regression
Residual

JD
BKN
JMN
BTH

Mean square
4427.62
3488.81
189.155
2397.94
20 985.8
2694.96
3614.83
2323.96
1824.54
5097.64

Regression equation

d.f.

F-ratio

P -value

Y = 145.155 0.8647X

1
33
1
29
1
34
1
31
1
31

1.27

0.268

0.08

0.780

7.79

0.008

1.56

0.2217

0.36

0.5540

Y = 126.537 0.1788X
Y = 173.792 2.0369X
Y = 141.675 0.6811X
Y = 187.838 0.6894X

LAB, laboratory; JD, Jodhpur; BKN, Bikaner; JMN, Jamnagar; BTH, Bathinda.
Laboratory strain LAB

Jodhpur strain JD

240

Y = 145.155 -0.8647X

200

Eggs/egg raft

Eggs/egg raft

240

160
120
80
40

Y=126.537 -0.1788X

200
160
120
80
40

0
0

10

20

30

40

50

60

0
Age of female, Days

(A)

30

40

50

60

Age of female, Days

240

300
Y =173.792 - 2.0369X

250

Eggs/egg raft

Eggs/egg raft

20

Jamnagar strain JMN

Bikaner strain BKN

200
150
100

Y = 141.675-0.6811X

200
160
120
80
40

50

0
0
(C)

10

(B)

10

20

30

40

50

Age of female, Days

20

40

60

80

Age of female, Days

(D)
Bathinda strain BTH

Eggs/egg raft

300

Y = 187.83- 0.6864X

250
200
150
100
50
0
0

(E)

10

20

30

40

50

Age of female, Days

Fig. 6. Regression line between eggs per egg raft and age of female (days) in Culex quinquefasciatus in (A) LAB, (B) JD, (C) BKN, (D) JMN
and (E) BTH strains.
2011 The Authors
Medical and Veterinary Entomology 2011 The Royal Entomological Society, Medical and Veterinary Entomology, doi: 10.1111/j.1365-2915.2010.00939.x

Net reproduction rate

Culex quinquefasciatus life tables 9

female life expectancy and R0 (0.850.91), but relatively
weakly with rm , and T (0.460.59).

180
160
140
120
100
80
60
40
20
0

Comparison of r-strategy among the different strains

Among the different field strains, JMN had the longest

female survival period, DT and the lowest rm and in
comparison with BTH, BKN and JD, indicating that JMN
was superior in r-strategy. LAB appeared to be a weaker
r-strategist than all the field strains because it required the
longest DT and had the least rm and .

11 16 21 26 31 36 41 46 51 56 61
Days

Lab

JD

BKN

JMN

BTH

Fig. 7. Cumulative net reproductive rate (R0 ) in Culex quinquefasciatus in the LAB, JD, BKN, JMN and BTH strains.

JD and BKN and between BTH and JMN, but no significant

differences between JD and BTH or between BKN and JMN.
The mean generation time (T ) was longest in LAB and shortest
in BTH. The LAB strain differed significantly from JD and
BTH for T , but not from BKN and JMN. The longest doubling
time (DT ) was observed in LAB, followed by JMN, BKN, JD
and BTH, but the differences between strains was not always
significant (Table 6).

Cluster analysis
Analysis of all Cx. quinquefasciatus strains indicated two
clusters, consisting of BKN and LAB, and JD, BTH and
JMN, respectively (Fig. 8). The distance between the clusters was 22.20. Cluster analysis also indicated a closer
relationship between JD and BTH than between JMN
and BKN.

Correlation analysis between environmental factors

and life-table attributes
Table 7 indicates that annual temperature ranges in the four
field locations were strongly correlated with egg production,
female life expectancy, R0 , rm , and T (0.770.97). Mean
annual rainfall also correlated strongly with egg production,

Discussion
The life-table parameters and other biological attributes in
five strains of Cx. quinquefasciatus originating from different
geographical areas varied with ecological conditions.
Apparent and actual mortality in eggs was found to range
from 4.4% (JD and BTH) to 19.5% (BKN); values in the other
two strains were relatively close to that in BKN. Rates of actual
mortality of larval stages ranged from 6.6% (JMN) to 32.3%
(BKN), with values for the other strains falling between these
extremes. The cumulative effect of mortalities of eggs, larvae
and pupae resulted in lower rates of adult emergence, ranging
from 42.0% (BKN) to 74.0% (BTH); values for the other
strains fell within 10% of that for BTH. The high mortality in
BKN may be associated with its habitat, which is very prone
to drought and is almost completely characterized by desert
conditions. By contrast, JMN has a significantly higher rate of
adult emergence than BKN despite a non-significant difference
in egg hatching because the conditions in which it lives provide
better support for larval and pupal development. In other
studies, rates of adult emergence were low for four strains of
Anopheles sacharovi Favr (Diptera: Culicidae) (11.414.6%)
(Yurttas & Alten, 2006), but exceeded 50% from first instars
of field-collected Anopheles gambiae Giles s.s. from Burkina
Faso (Lehmann et al., 2006). These studies revealed that
variations in immature mosquito developmental rates depend
on habitat conditions.
The percentage of eggs hatching has been reported to
be 82.1% in Culex pipiens fatigans (synonymous with
Cx. quinquefasciatus; Gomez et al., 1977) and 79.6% in

Table 6. Comparison of adult life-table attributes in Culex quinquefasciatus of four strains from different geographical areas and a strain raised
under laboratory conditions.

Parameters

LAB strain,
mean SE

JD strain,
mean SE

BKN strain,
mean SE

JMN strain,
mean SE

BTH strain,
mean SE

LSD

R0
rm

T
DT

107.6
0.15 0.00a
1.16 0.00a
31.6 0.80a
4.7 0.11a

147.4
0.20 0.00b
1.23 0.00b
24.5 0.38b
3.4 0.05b

122.9
0.17 0.00c
1.18 0.00a
28.8 0.70a
4.1 0.05c

162.2
0.16 0.01ac
1.17 0.01a
31.4 2.08a
4.3 0.28c

135.6
0.21 0.02b
1.24 0.02b
22.9 1.67b
3.2 0.24b

8.76
0.02
0.02
2.94
0.24

1.96a

2.83b

7.89c

1.96d

1.29e

Mean values within rows indicated by same letter do not differ significantly.
LAB, laboratory; JD, Jodhpur; BKN, Bikaner; JMN, Jamnagar; BTH, Bathinda; SE, standard error; LSD, least significant difference; R0 , net
reproductive rate; rm , intrinsic rate of increase; , finite rate of increase; T , mean generation time; DT, doubling time.
2011 The Authors
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10 D. S. Suman et al.
24

Distance

20
16
12
8
4
JMN

BTH

JD

BKN

Lab

Fig. 8. Dendrogram of the four field strains (JD, BKN, JMN and
BTH) and a laboratory strain (LAB) of Culex quinquefasciatus
showing similarities in various life-table attributes. The distance
coefficient is measured using a squared Euclidean method.
Table 7. Correlation analysis of various life-table and biological
attributes for the JD, BKN, JMN and BTH strains of Culex
quinquefasciatus for temperature range and mean annual rainfall.
Correlation coefficient (r )
Attributes

Temperature range

Mean annual rainfall

Egg production/cohort
Female life expectancy
R0
rm

0.79
0.97
0.76
0.82
0.77
0.87

0.90
0.86
0.91
0.51
0.46
0.59

R0 , net reproductive rate; rm , intrinsic rate of increase; , finite rate of

increase; T , mean generation time.

Cx. quinquefasciatus (Suleman & Reisen, 1979), which are

close to the values reported in the present study (Table 1).
Variations in percentages of eggs hatching have also been
recorded in Cx. tritaeniorhynchus, for which higher rates
emerged in Indian subcontinental strains (93.0%) than in
a Japanese strain (78.8%) (Reisen et al., 1979). However,
significantly lower egg-hatching rates were recorded in four
strains of An. sacharovi from southeastern Anatolia, Turkey,
leading to high overall real mortality rates (85.488.6%;
Yurttas & Alten, 2006). The eggs of strains collected
in the present study from the most arid environments
(BKN, JD) required a shorter hatching period than those
of semi-arid strains (JMN, BTH), which suggests that short
hatching times may be a positive factor for survival in arid
conditions.
In the present study, the mean development time from egg
to adult was found to vary little, with four of the strains
showing a range of <1 day (11.612.2 days in LAB, BKN,
JMN and BTH). Only the JD strain differed significantly
from these by taking an extra 2 days (14.8 days). These values were all shorter than the developmental period reported
for Culex tarsalis Coquillett collected in California, U.S.A.
(15.019.8 days; Reisen & Milby, 1987) and maintained in
laboratory colonies for several years. These longer development times may reflect rearing at a lower temperature

(25 C). Values similar to those reported here were found for
An. sacharovi from southeastern Anatolia, Turkey (Yurttas &
Alten, 2006).
Rates of immature survival to adult emergence ranged from
42.0% (BKN) to 74.0% (BTH), with rates in the three other
strains (LAB, JD and JMN) falling within 10% of the highest value. It is interesting to note that, although BKN had the
lowest rate of adult emergence, its female : male ratio was significantly higher than that in any of the other strains (1.25 vs.
1.10), whereas BTH had the highest rate of adult emergence
and the lowest female : male ratio (0.98). This indicates that
a low rate of adult emergence may be compensated for by a
high ratio of female emergence.
Life expectancy is an important attribute of mosquito populations in relation to their survival and likelihood of transmitting pathogens. Studies on vector transmission indicate
that microfilariae require 1617 days in the mosquito vector
to reach the infective stage (Bancroft, 1899) and the period
of microfilariae development varies with season (Chandra
et al., 1997), decreasing in duration at high temperatures and
high humidities to complete the cycle of development in
1014 days, but increasing to up to 42 days in low temperatures (Manson-Bahr & Bell, 2003). The present study found
life expectancy to be longer in females (25.236.9 days) than
in males (18.221.3 days) in all strains. Similarly, Walter &
Hacker (1974), Gomez et al. (1977) and Suleman & Reisen
(1979) recorded longer life expectancies in females than in
males in different strains of Cx. quinquefasciatus. The males
of JD, BKN and JMN had significantly shorter lifespans than
those of BTH and LAB. However, females of JD and JMN
lived significantly longer than those of LAB, BTH and BKN.
Similar variabilities among strains were observed by Walter
& Hacker (1974) in Cx. quinquefasciatus from Bangkok, Vero
Beach and Houston. In the present study, the mortality patterns
of different strains of Cx. quinquefasciatus exhibited approximate Slobodkin type II curves that were similar to the mortality
patterns of different strains of Phlebotomus papatasi (Scopoli)
(Diptera: Psychodidae) and were characterized by relatively
constant mortality rates throughout all life stages (Belen &
Alten, 2006).
No significant differences among the strains were apparent
for pre- and post-oviposition periods (Table 2). The oviposition
period in JD, however, differed significantly from those in the
other strains and was 3 days shorter than the others, with a
mean of only 21 days. Significant differences in oviposition
period were also reported in various strains of An. sacharovi
(Yurttas & Alten, 2006).
Significant differences among all the strains emerged in
the various measures of fecundity, including total egg production/cohort (range: 10 76316 218), eggs/female lifespan
(range: 512.4812.3) and eggs/female/day (range: 20.530.8),
except that mean numbers of eggs/female/day were similar in LAB and BKN, and in JD and JMN. Reisen et al.
(1984) recorded fewer eggs/female/day (range: 6.320.7) in
Cx. tarsalis. Yurttas & Alten (2006) also found variations in
egg production/female in four geographically distinct strains
of An. sacharovi.

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Culex quinquefasciatus life tables 11

Numbers of eggs/raft varied significantly among strains of
Cx. quinquefasciatus (Table 2), ranging from 161.1 (JMN) to
207.5 (BTH), whereas the mean number of rafts/cohort was
nearly opposite, ranging from 65.3 (BTH) to 100.7 (JMN). This
is not surprising because high numbers of eggs/raft probably
reflect the production of relatively few rafts/female and vice
versa. The construction of large egg rafts may consume more
reproductive energy and time than the construction of smaller
egg rafts and thus females that laid small egg rafts were
able to produce more eggs over their lifetime. Significant
differences in mean numbers of eggs/raft were observed among
all four field-collected strains, but values in BKN and JMN
did not differ significantly from that in LAB. Variations in
the mean number of eggs/raft among different strains of Culex
mosquitoes have been reported previously (Walter & Hacker,
1974; Reisen et al., 1979, 1984).
Interestingly, the strains from areas with the greatest
range of annual temperatures (Bathinda, Bikaner and Jodhpur) face tougher conditions for survival in nature and these
mosquitoes typically had a shorter female life expectancy
(25.230.8 days), laid eggs on most days (21.026.3 days) and
constructed larger egg rafts (170.2207.5 eggs/raft), indicating
that these strains invest the most energy in rapid reproduction.
By contrast, the Jamnagar strain, which comes from the area
with the smallest range of annual temperatures and hence more
optimal conditions for mosquitoes, had the longest female life
expectancy (mean: 36.9 days), but a shorter oviposition period
(mean: 24.7 days) and laid fewer eggs/raft (mean: 161.1). Fox
& Czesak (2000) reviewed the evolutionary ecology of progeny
size and concluded that more hostile environmental conditions
may favour the production of a greater number of offspring
in order to increase the probability that some will survive to
adulthood. Thus, individuals from unfavourable environments
are inherently committed to put maximum effort into the production of more offspring to ensure that an adequate number
survive to reproduce in their turn.
In the present investigation, the mean number of egg
rafts/female (Fig. 4, Table 3) and the mean number of
eggs/female (Fig. 5, Table 4) increased with female age. The
mean number of eggs/raft, however, decreased with female
age (Fig. 6, Table 5). These observations indicate that cumulative fecundity increased as a function of female age as
a result of an increase in the frequency of the deposition
of progressively smaller rafts. Similarly, Walter & Hacker
(1974) found the number of eggs/raft to decrease with increasing age in a range of geographically distinct strains of
Cx. quinquefasciatus. Suleman & Reisen (1979) recorded similar observations in Cx. quinquefasciatus and Reisen et al.
(1984) did so in Cx. tarsalis.
Species with relatively high net reproductive rates (R0 ) tend
to have a competitive advantage over sympatric species that
occupy similar ecological niches. Gubler (1970) observed the
competitive displacement of Aedes polynesiensis Marks by
Aedes albopictus (Skuse) (Diptera: Culicidae), and showed
that the latter species was superior in female survival and net
reproductive rate. In the present investigations, the R0 values in
all wild strains (122.9162.2) were significantly greater than in
LAB (107.6) and differed significantly amongst one amother,
which suggests that varying degrees of fitness of the wild

strains may have been imposed by different levels of selection

pressure on the reproductive rate for each strain. Similarly,
significant differences in R0 for Cx. quinquefasciatus have
been reported in strains from Bangkok (8.1), Vero Beach (24.3)
and Houston (37.7) (Walter & Hacker, 1974), from Caracas,
Venezuela (80.7) (Gomez et al., 1977) and from Peshawar,
Pakistan (58.6) (Suleman & Reisen, 1979); however these
values are all lower than those found in the present study.
Reisen et al. (1979) observed variations inR0 in nine strains
of Cx. tritaeniorhynchus, with the highest values reported in
strains from Karachi (98.5) and Taipei (84.9), and the lowest
reported in three strains from Japan (14.431.2). The present
and referenced data show a consistent trend for mosquito
populations in colder climates to have lowerR0 values than
populations in warmer climates. Yurttas & Alten (2006) also
found significant differences in R0 (range: 3.212.8) among
different strains of An. sacharovi, which are again, lower than
those in the present study.
In the current study, differences in rm (0.150.21),
(1.161.24) and DT (3.234.67) were observed among
the various strains of Cx. quinquefasciatus and some of
these differences were significant. Similarly, Walter &
Hacker (1974) found significant differences in rm between
a Cx. quinquefasciatus strain from Bangkok and two other
strains (Vero Beach and Houston), although the latter two
strains did not differ significantly in rm from each other.
The rm values in LAB, BKN and JMN are closest to that
in the Caracas strain of Cx. p. fatigans (Gomez et al., 1977),
but much higher than the value in the Peshawar strain of
Cx. quinquefasciatus (Suleman & Reisen, 1979). Reisen et al.
(1979) reported greater differences in rm among various strains
of Cx. tritaeniorhynchus from the Indo-Pakistani subcontinent
(0.160.19) and Taiwan (0.18) than among Japanese strains
(0.080.12). This appears to reflect a greater need on the part
of the Japanese strains for a longer mean generation period
(28.229.8 days) than the Indo-Pakistani (22.224.1 days) and
Taiwanese (24.0 days) strains. A comparison of these data
indicates that warmer climates have a positive effect on rm ,
increasing inherent capacity for reproduction.
In the present study, the mean generation time (T ) ranged
from 22.9 days to 31.6 days among the various strains of
Cx. quinquefasciatus. Higher values for T (39.6 days and
44.7 days) were recorded by Gomez et al. (1977) and Suleman
& Reisen (1979) and lower values by Walter & Hacker (1974)
in the Bangkok (32.1), Vero Beach (26.8) and Houston (28.5)
strains of Cx. quinquefasciatus.
The cluster analysis of various life-table and biological
attributes of the four wild strains of Cx. quinquefasciatus
revealed that JD was biologically closer to BTH than to BKN
and JMN, which may reflect cumulative differences in their
respective ecologies.
Although Bathinda (BTH) and Jamnagar (JMN) are 1000
km apart, they both lie in the same semi-arid zone. Average
annual rainfall is greater in Jamnagar than in Bathinda and RH
is also greater for most of the year in Jamnagar as a result
of its coastal location. The range of annual temperatures is
narrower for Jamnagar than Bathinda. Accordingly, we might
expect that environmental conditions in Jamnagar would favour
survival and reproductive potential and would support a shorter

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12 D. S. Suman et al.
egg-hatching period (1.96 days), longer female life expectancy
(36.9 days), a higher net reproductive rate (162.2) and a longer
mean generation time (31.4 days) than conditions in BTH
(2.42 days, 25.2 days, 135.6 days and 22.9 days, respectively).
Jodhpur and Bikaner both lie in arid zones. Although Jodhpur has relatively higher rainfall than Bikaner, these sites
would be expected to be less favourable than Jamnagar and
Bathinda for mosquito survival and reproduction. The positive
correlations between rainfall and various life-table parameters for the wild strains indicate that environmental factors
do indeed have an important evolutionary effect on mosquito
life strategies that can be detected even after several generations of colonization. The JD, BKN and BTH strains of
Cx. quinquefasciatus have relatively low DT as a result of
higher rm and , which perhaps reflects adaptations to more
extreme environmental conditions than occur in Jamnagar,
whereas the JMN strain had lower rm and values and higher
DT, despite high R0 and T values. By contrast, the LAB strain
had a relatively high DT value as a result of having the lowest R0 , rm and in comparison with the four wild strains.
Crovello & Hacker (1972) also reported variation in the reproductive potential of 13 strains of Aedes aegypti L. from a range
of habitats and found forest strains to show the lowest reproductive potential as a result of more homogeneous levels of
environmental stress than found in urban and semi-urban areas,
where there is a greater degree of fluctuation in environmental
conditions. The evolution of an r-strategy has been described
for Cx. quinquefasciatus by Rabinovich (1974) and supported
by Gomez et al. (1977) and Suleman & Reisen (1979). Reisen
et al. (1979) also showed that strains of Cx. tritaeniorhynchus
from Taiwan and the Indo-Pakistan subcontinent were more
r-selected as a result of the more variable ecological systems in which they live compared with Japanese strains, which
originated from more stable, cooler and more humid maritime
climates. These strains of Cx. tritaeniorhynchus were lesser rstrategists than domestic species, such as Cx. quinquefasciatus
and Ae. aegypti (Schlosser & Buffington, 1977). Analysis of
the various parameters in the present study indicates that,
among the wild strains, BTH, BKN and JD are more efficient
r-strategists as they face more difficult environmental conditions than does JMN, which exists in more homogeneous
environmental conditions. Interestingly, Suman et al. (2009)
showed the JMN strain to differ from the JD, BKN and BTH
strains of this mosquito for various egg-related attributes. Furthermore, higher R0 , rm and values and lower T and DT
values were observed in the wild strains than in LAB, which is
consistent with the hypothesis that harsher environmental conditions select for high r-strategists in the field. Strains with high
r-strategies may be more difficult to control because, when a
population is diminished, it is able to re-establish more quickly
than a less able r-strategist.
The results of this study extend our knowledge of the impact
of the environment on a wide range of life history traits in local
populations of disease vectors. The study also provides detailed
information which is valuable for increasing the accuracy of
the population dynamic models that are an essential component
of the decision-making tools used to plan and monitor vector
control programmes (World Health Organization, 2004).

Acknowledgements
We thank Dr R Vijayaraghavan, Director, Defence Research
and Development Establishment, Gwalior, for his support and
interest in this work.

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Medical and Veterinary Entomology 2011 The Royal Entomological Society, Medical and Veterinary Entomology, doi: 10.1111/j.1365-2915.2010.00939.x